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Title Occurrence of the European Ascidian Ascidiella Scabra

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Title Occurrence of the European Ascidian Ascidiella Scabra View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Kyoto University Research Information Repository Occurrence of the European ascidian Ascidiella scabra (Müller, Title 1776) in the 19 century in Nagasaki, Japan, probably as an ephemeral alien species Author(s) NISHIKAWA, Teruaki; OTANI, Michio Contributions from the Biological Laboratory, Kyoto Citation University (2004), 29(4): 401-408 Issue Date 2004-07-21 URL http://hdl.handle.net/2433/156412 Right Type Departmental Bulletin Paper Textversion publisher Kyoto University Contr. biol. lab. Kyoto Univ., Vol. 29, pp. 401ZK)8. Issued 21 July 2004 Occurrence of the European ascidian Ascidiella scabra (Milller, 1776) in the 19 century in Nagasaki, Japan, probably as an ephemeral alien species Teruaki NISHIKAWA' and Michio OTANI2 Thei Nagoya University Museum, Chikusa-ku, Nagoya, 464-8601 Japan 2Marine Ecological Institute Inc., 3-3-4 Harada Moto-machi, Toyonaka, 561 -0808 Japan ABSTRACT The generic affiliation of the single Japanese record of the ascidian genus Ascidietta, as "A. virginea (MtiIIer)" by Hartmeyer in 1902, was questioned by Nishikawa in 1995 because of the genus's natura1 distribution being confined to Atlantic waters, the somewhat confused history of taxonomy in the genus and allies, and of the insufficient description. Our personal reexamination of the material collected from Nagasaki and deposited in the Museum fUr Naturkunde der Humboldt-Universitat zu Berlin reyeals that it is identical to Ascidietla scabra (Mti11er, 1776), so far recorded exclusively from European boreal to warm-temperate waters. Combining imperfect label information of the material with historical knowledge, the material is supposed to have been collected in February, 1861, by Mr. Otto SchottmUller, a member of the Prussian East Asia Expedition led by Graf zu Eulenburg. It is proposed that the material probably represents an ephemeral population of an alien species. KEY WORDS Ascidielta scabra / ascidian 1 alien species X ephemeral population 1 Japan Introduction More and more attention has come to be paid to the so-called "alien or inyasive species issues" in terms of conservation of biodiversity (see, e.g., Ecological Society of Japan, 2002). When one confronts the issues, the problem is that it is often not easy to decide precisely whether a local population is indigenous or not. This is partly because the biota change through time has been insufficiently documented, and partly because the specific identification tends to become unstable due to subsequent ineyitable taxonomic changes (caused, e.g., by discovery of cryptic species). It follows that the actual specimens that have been maintained for many years in museums can play a significant role, though more or less limited according to ways and conditions of preservation, as the "liying" evidence to be used for reconstruction of the past biota. The genus Ascidiella in the Japanese ascidian fauna has been represented by a single isolated record of "Ascidiella virginea (MUIIer)" in Hartmeyer (1906, p. 22). His paper is one of the first two monographic works on it, with the other by the first Japanese ascidian taxonomist Prof. Oka Asajiro (see Oka, 1906). The specific name virginea is now combined with the generic name Ascidia, as clearly distinct from Ascidiella, thus used as a valid species name Ascidia virginea Mti11er, 1776. Considering the taxonomic and nomenclatural confusion in the distinction between these two genera and in the specific identification 402 T. NISHIKAWA & M. OTANI among their congeners in the past, together with Hartmeyer's insufficient description, Nishikawa (1995) once suggested that Hartmeyer's record might represent the occurrence of a species of the genus Ascidia, whose many species occur in Japanese waters (see, e.g., Tokioka, 1963; Nishikawa, 1995). On the other hand, many previous records of Ascidietla virginea (MUIIer, 1776), excepting the Japanese one, were incorporated in those of Ascidietla aspersa (MUIIer, 1776) by Hartmeyer (1924); this species has been collected mainly from European waters, and therefore, is believed to have been introduced to New Zealand since the early 1900s (Cranfield et al., 1998) and to India since 1976 (Nagabhushanam and Krishnamoorthy, 1992). Therefore, another hypothesis is also possible that Hartmeyer's record represents an ephemeral occurrence of Ascidiella aspersa in Japanese waters. To test these hypotheses, Hartmeyer's specimens long kept in the Museum fUr Naturkunde der Humboldt-Universittit zu Berlin was fortunately reexamined. As a result, both of the hypotheses are rejected, although the specimens can still be regarded to represent an ephemeral Japanese population of another alien species. Historical Background Concerning Ascidiella scabra and ARIies The genus Ascidielta is unique among ascidians of the family Ascidiidae in having no secondary branchial papillae on the internal longitudinal vessels (see, e.g., Beni11, 1950). After a long history of taxonomic and nomenclatural confusion, the genus is now considered to consist of the three species, A. aspersa (Mtiller, 1776), A. scabra (MUIIer, 1776), and A. senegalensis Michaelsen, 1914. 0f these, A. senegalensis, so far recorded rarely from the Atlantic coasts of northwestern Africa, is peculiar in having the hook-like anterior ends of inner longitudinal vessels (see, e.g., Michaelsen, 1914; Monniot, 1969). A. aspersa and A, scabra were once considered conspecific by some authors including the influential work by Hartmeyer (1924). However, according to a historical review by Berrill (1950), the conspecificity was doubted by some researchers on the basis of differences between them in the body size and shape (especially in the position of atrial siphon), the minimal body size when matured, the size of outer follicle cells of eggs, and in that "in A. aspersa the number of longitudinal vessels always outnumbers that of the tentacles, whereas in A. scabra the reverse is true" (p. 157). These differences came mainly from a detailed morphological comparison between the two species given by Lindsay and Thompson (1930). The number of longitudinal vessels mentioned is doubtlessly that of the vessels on each side, instead of their total number, because "the Iongitudinal bars [=vessels] were enumerated in a transverse direction in the right half of the branchial sac" (Lindsay and Thompson, 1930, p. 11). Subsequently, Millar (1966, p. 57) gave a taxonomic key for the identification of Scandinayian ascidians to distinguish one from the other, such as: A. aspersa is characterized by the "body length often exceeds 6 cm; tentacles fewer than internal longitudinal bars [=inner longitudinal vessels on each side]", while A. scabra by the "body length usually under 4 cm; tentacles more numerous than internal longitudinal European ascidian occurring in the I9 century in Nagasaki 403 bars". In contrast to A. aspersa, A. scabra has not yet been recorded from outside European waters, so far as we are aware. Hartmeyer's (1906) Japanese record of "Ascidiella virginea (MUIIer)", which is the main topic of the present paper, was based on a certain number of specimens collected "von Schottmtiller bei Nagasaki", as he detected from old label information, and deposited in the "Berliner Museum" (p. 22). At the same time, Hartmeyer reserved the decisive specific identification of the specimens, taking the possibility of mislabeling into account. Later, Hartmeyer (1924) completely neglected to mention the record from Japan, excluding it from the geographical distribution of Ascidiella aspersa (sensu lato, see aboye), compiled from a thorough survey of the previous records which contained those once referred to as A. virginea; howeyer, according to the catalogue of the museum he finally identified the Japanese material as A. aspersa (see below), In the present study, the label information for the material is to be considered reliable and correct, rather than false due to mislabeling. Needless to mention, the possibility of mislabeling should always be kept in mind when one treats museum specimens, but it may not be very wise to adopt this possibility primarily. In this case, Hartmeyer seems to have ruled out such a possibility for the Japanese population to be transported from Europe by human movements, as will be discussed later. In his days, it seems that a species' potentiality of introduction into distant districts by ships was not accepted unanimously, as may be imagined from Visscher's (1938, p. 342) statement "that there is no evidence of distribution of barnacles from one port to another by virtue of the attachment on bottoms of ships". Materials and Methods The material examined was labeled "ZMB 1051, Ascidiella aspersa (MUIIer), Nagasaki, Schottmif11er", kept in the Museum ftir Naturkunde der Humboldt-Universitat zu Berlin, consisting of 18 individuals; a 44.5 mm long one was dissected with permission. The mentioned label information is quite the same as that seen in the museum's catalogue of "Protochordata", beautifully hand-written (probably by Hartmeyer himselb as "1051, Ascidiella aspersa (Mti11er), Nagasaki, Schottmti11er S. [=coll.l 1 A. virginea, Hartmeyer in: Zool. Anz., v. 31, p. 22, 1906 [=Hartmeyer's (1906) paper]". Therefore, it seems certain that the material was ultimately identified as A. aspersa by Hartmeyer, as an unpublished taxonomic decision (see above). We haye tried to collect further specimens of Ascidiella in Nagasaki Bay, and also to find any old specimens collected there in seyeral museums and institutions in Japan, including the Oka collection of ascidians in the University of Tsukuba as the most probable candidate, but in vain. Moreover, a literature survey of the ascidian fauna around Nagasaki was performed, but we could not detect any positive evidence for the existence of Ascidietta, other than Hartmeyer's (1906). 404 T. NISHIKAWA&M. OTANI Taxonomic Results Ascidiella scabra (Mti11er, 1776) (Fig. 1) Limited Synonymy and Previous Records: Ascidia scabra Mti11er, 1776, p. 225 Ascidiella scabra: Berrill, 1950, pp. 158-159, fig. 48; Millar, 1952, pp. 23-24; 1959, p. 4; 1966, p. 58, fig. 38; Turon, 1987, pp. 139-140. Ascidielta aspersa var.
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